增强的UV-B辐射对植物影响的研究

综述了国内外有关UV－B辐射对植物影响的研究现状与动态,讨论了增强的UV－B辐射对植物生长及形态结构、植物生理生化代谢、植物遗传物质、UV－B吸收物质及某些基因表达和种群及生态系统的影响。展望了增强的UV－B辐射对植物影响领域中值得深入探讨的问题。     

UV-B辐射对植物花粉萌发率和花粉管生长的累积效应

研究了19种植物花粉在不同UV－B辐射强度和辐照时间下其萌发率和花粉管伸长的变化,结果表明,UV－B辐射增加显著抑制大多数植物花粉的萌发率和花粉管生长;与对照相比,较高强度的UV－B对花粉的抑制作用大于较低强度;几个种的花粉萌发率及花粉管生长对UV－B增强不敏感,甚至被UV－B辐射所促进;辐射时间越长,对花粉抑制作用愈大,说明具有辐射累积效应,由此可知,植物花粉的萌发过程对UV－B的敏感性变化在自然条件下将会产生严重的生态学后果。     

增强UV—B辐射对高山植物麻花艽净光合速率的影响

在高寒矮嵩草（Kobresia humilis)草甸地区以太阳短波辐射为背景,建立了人工增强UV－B辐射的实验装置,每天增补15．8kJ.m^-2的辐射剂量,模拟平流层臭氧破坏约5％时近地表面太阳UV－B辐射的增强。观测表明：UV－B辐射的增强对麻花艽（Gentiana stramianea)植物的光合作用无明显的抑制或伤害作用。相反,在早晨补充UV－B辐射的短时间内,叶片的Pn随Gs的增大而有所提高。随着UV－B辐射时间的延长,在11：30－12：30,Pn和Gs有所降低。UV－B辐射时间进一步延长后（约14：00以后）,处理和对照组叶片Pn和Gs的差异趋向不明显,增强太阳UV－B辐射后,麻花艽叶片的光合色素并无明显变化,UV－B吸收物质的含量无明显变化,麻花艽叶片厚度的直接测量表明,增强UV－B辐射能明显提高叶片的厚度。叶片厚度的增加可补偿增强UV－B辐射辐后引起的光合色素的光降解,改善单位叶面积为基础的光合速率,是高原植物对强UV－B辐射的一种适应方式。     

紫外线B辐射与植物体内酚类次生代谢的关系

概述了植物酚类次生代谢产物对紫外线B(UV—B)增强引起的辐射伤害的防御作用,酚类次生代谢与其它防护机制的相互关系。     

增强UV-B辐射与干旱复合处理对小麦幼苗生理特性的影响

为研究由于平流层臭氧层减薄紫外线B辐射增强在干旱地区对春小麦生理特性的影响的特殊性,模拟平流层臭氧减少20％时辐射到地表的紫外线B（UV－B,280－315nm)的增强和水分胁迫（－0.5Mpa,聚乙二醇PEG－6000处理获得）,通过测定两种胁迫下春小麦（Triticum aestivum L.)叶绿素含量、类黄酮含量、水势、细胞膜相对透性、超氧歧化酶（SOD）活性及丙二醛（MDA）含量,研究了增强UV－B辐射和水分胁迫复合作用对温室种植的小麦幼苗生理生化的影响。实验结果表明,虽然水分胁迫和UV－B辐射单独或复合处理都使春小麦的叶绿素含量降低,但仅UV－B辐射增强单独处理显著地降低小麦叶绿素a、b和总叶绿素的含量,而水分胁迫以及复合处理对叶绿素的含量的降低作用不显著。两种胁迫无论是单独作用还是复合作用均能使类黄酮含量升高,并且处理第3天比第1天高出近50％,复合处理下类黄酮的含量大于两个因子单独处理。UV－B辐射和水分胁迫处理1d对春小麦叶片的相对电导率的影响不明显,处理3d后两种胁迫下相对电导率均上升,表明膜透性增加,其中水分胁迫作用下增加尤其明显。膜质过氧化产物丙二醛的含量,在两种因子单独和复合作用下都升高,说明膜的生理功能受到了一定的不利影响。活性氧清除剂超氧化物歧化酶（SOD）的活性在各种处理下,都没有发生改变。虽然增强的UV－B辐射和干旱处理一样,会显著降低植物水势,但是,UV－B辐射与干旱同时处理时叶片水势降低的程度,不但没有比两者分别处理时降低程度之和低,而且比单做干旱处理时的降低程度还低,这表明UV－B辐射和干旱胁迫同时处理时,UV－B辐射不但没有加重水分胁迫,反而减轻了干旱对春小麦生长的胁迫。由此可以认为,在干旱条件下,增强UV－B辐射不会加剧而是有利于提高小麦对干旱的抗性。     

大豆作物响应增强UV—B辐射的品种差异

田间条件下模拟20％平流层臭氧层衰减,紫外线（UV－B,280－315nm)辐射增强,研究了UV－B对2个大豆（Glycin max (L.)Merr.)品种黑豆和晋豆生长,光合作用和稳定碳同位素组成的影响,结果表明,晋豆比黑豆对UV－B有较强的抗性或不敏感,表现为增强的UV－B辐射显著抑制黑豆的生长和株高,叶、茎、根和总生物量以及株高全部降低,而晋豆仅茎重和株高降低;晋豆的色素含量（叶绿素a,b,类胡萝卜素和类黄酮）不受UV－B辐射影响,在UV－B辐射下黑豆的净光合作用,气孔导度,胞间CO2浓度和蒸腾作用以及不分利用效率明显下降,而晋豆只有气孔导度和蒸腾作用减少,这可能与晋豆本身含有较高的类黄酮及较多的表皮毛和遗传特性有关,用叶片稳定碳同位素组成（δ^13C值）的分析也证明晋豆对UV－B辐射不敏感,由此看来,大豆品种对UV－B辐射的反应差异可以通过δ^13C值来判定。     

重金属镉（Cd）和增强UV—B辐射复合对大豆生长和生理代谢的影响

研究了大豆的生长、生物量、抗氧化酶活性和吲哚乙酸（IAA）氧化酶活性在Cd^2 、UV－B辐射和二者复合胁迫（Cd UV-B)下的变化。结果表明,Cd^2 和UV－B辐射都抑制大豆生长,并显著抑制根的伸长,二者复合后加强了对根伸长的抑制。UV－B辐射显著增强了POD、SOD活性,Cd^2 对POD活性影响不明显,但却拮抗UV－B对POD活性的诱导,SOD活性在各种胁迫下显著增强。虽然Cd%2 对叶片类黄酮含量影响不明显,但对UV－B诱导的类黄酮合成有一定影响。IAA氧化酶活性在复合作用下下降,可能是复合胁迫影响大豆生长的重要因素之一。     

UV-B辐射对香蕉光合作用和不同氮源利用的影响

生长在NO3^--N、NH4^--N和NH4NO3－N的香蕉叶片有相近似的最大光合速率,UV－B辐射引起生长在不同氮源的香蕉叶片光合速率、表现量子产率和光肥利用效率的降低。UV－B辐射使生长在不同氮源的植株叶面积干重和叶氮含是降低。生长在NH4^--N的植株Vcmax和Jmax均较生长在其它氮源的高。UV－B辐射引起生长在NH4^-N的植株Vcmax和Jmax降低较相同处理的NO3^-－N和NH4NO3－N植株明显,表明生长在NH4^ －N的香蕉对UV－B辐射更加敏感。UV－B辐射改变植株的叶片的碳氢比和碳氮比。经过UV－B辐射处理的NH4^ －N生长植株的碳氮生长在NO3^--N和NH4NO3－N的低。UV－B辐射可能改变植株对不同氮源的吸收利用,从而引起碳氮代谢和酸碱调节的变化。UV－B辐射降低叶氮在Rubisco和生物力能学组分的分配系数,可能使这些组分合成减少,使叶片光调节的变化。UV－B辐射降低叶氮在Rubisco和生物力能学组分的分配系数,可能使这些组分合成减少,使叶片光合速率下降。结果表明,生长在不同氮源的香蕉植树对UV－B辐射有不同响应,NH4^ -N有利于主要光合参数增高,但其对UV－B辐射亦最为敏感。氮供应受限制或植株生长在中性盐如NH4NO3－N则对UV－B辐射不甚敏感。     

田间增强UV—B辐射对麦田生态系统K营养和累积的影响

研究大田栽培和自然光条件下,模拟UV－B辐射（UV－B,280－315nm)增强对麦田生态系统K营养和累积的影响,UV－B辐射显著影响春小麦不同生育期各部位K含量和群体K累积,并显著降低群体K总累积,在5．31kJ.m^-2UV-B辐射下,春小麦群体K总累积的降低最显著,UV－B辐射降低春小麦群体K输出,标志着麦田生态系统K产投比降低,K循环功能下降,麦田土壤速效K含量增加是春小麦群体K输出降低的结果,并将导致土壤库中K储量的增加。     

增强紫外B辐射对植物及生态系统影响研究的发展趋势

介绍了一些有关紫外B辐射增强对植物及生态系统影响研究的新进展：1．许多研究已深入到分子水平;2．注意到对植物生长调控的研究;3．更加重视对植物防御、修复的研究;4．有关信号传导的研究日渐增多;5．对植物群体及生态系统影响的研究在不断扩大与加深;6．复合效应研究正在升温。推断今后在一段时间内,有关UV－B辐射对植物和生态系统影响的研究不但不会削弱,可能还会加强,特别分子水平的研究会大大增加,今后对群体和生态系统的研究会重视野外和长期效应的观测。我国在这一领域的研究起步晚,但近些年发展得较快,有部分研究已赶上国际研究进展的步伐。     

Changes in photosynthesis and antioxidant defenses of Picea asperata seedlings to enhanced ultraviolet-B and to nitrogen supply

The paper mainly studied the effects of ultraviolet‐B (UV‐B) radiation, nitrogen, and their combination on photosynthesis and antioxidant defenses of Picea asperata seedlings. The experimental design included two levels of UV‐B treatments (ambient UV‐B, 11.02 KJ m⁻² day⁻¹; enhanced UV‐B, 14.33 KJ m⁻² day⁻¹) and two nitrogen levels (0; 20 g m⁻² a⁻¹ N) – to determine whether the adverse effects of UV‐B are eased by supplemental nitrogen. Enhanced UV‐B significantly inhibited plant growth, net photosynthetic rate (A), stomatal conductance to water vapor (Gs), transpiration rate and photosynthetic pigment, and increased intercellular CO₂ concentration, UV‐B absorbing compounds, proline content, malondialdehyde (MDA) content, and activity of antioxidant enzymes (peroxidase (POD), superoxide dimutase, and glutathione reductase). Enhanced UV‐B also reduced needle DW and increased hydrogen peroxide (H₂O₂) content and the rate of superoxide radical (O₂⁻) production only under supplemental nitrogen. On the other hand, supplemental nitrogen increased plant growth, A, Gs, chlorophyll content and activity of antioxidant enzymes (POD, ascorbate peroxidase, and catalase), and reduced MDA content, H₂O₂ content, and the rate of O₂⁻ production only under ambient UV‐B, whereas supplemental nitrogen reduced activity of antioxidant enzymes under enhanced UV‐B. Carotenoids content, proline content, and UV‐B absorbing compounds increased under supplemental nitrogen. Moreover, significant UV‐B × nitrogen interaction was found on plant height, basal diameter, A, chlorophyll a, activity of antioxidant enzymes, H₂O₂, MDA, and proline content. These results implied that supplemental nitrogen was favorable for photosynthesis and antioxidant defenses of P.asperata seedlings under ambient UV‐B. However, supplemental nitrogen made the plants more sensitive to enhanced UV‐B, although some antioxidant indexes increased.     

Integration and scaling of UV‐B radiation effects on plants: from molecular interactions to whole plant responses

A process based model integrating the effects of UV‐B radiation to molecular level processes and their consequences to whole plant growth and development was developed from key parameters in the published literature. Model simulations showed that UV‐B radiation induced changes in plant metabolic and/or photosynthesis rates can result in plant growth inhibitions. The costs of effective epidermal UV‐B radiation absorptive compounds did not result in any significant changes in plant growth, but any associated metabolic costs effectively reduced the potential plant biomass. The model showed significant interactions between UV‐B radiation effects and temperature and any factor leading to inhibition of photosynthetic production or plant growth during the midday, but the effects were not cumulative for all factors. Vegetative growth were significantly delayed in species that do not exhibit reproductive cycles during a growing season, but vegetative growth and reproductive yield in species completing their life cycle in one growing season did not appear to be delayed more than 2–5 days, probably within the natural variability of the life cycles for many species. This is the first model to integrate the effects of increased UV‐B radiation through molecular level processes and their consequences to whole plant growth and development.     

Responses of photosynthetic properties and chloroplast ultrastructure of Bryum argenteum from a desert biological soil crust to elevated ultraviolet‐B radiation

Our understanding of plant responses to enhanced ultraviolet‐B (UV‐B) radiation has improved over recent decades. However, research on cryptogams is scarce and it remains controversial whether UV‐B radiation causes changes in physiology related to photosynthesis. To investigate the effects of supplementary UV‐B radiation on photosynthesis and chloroplast ultrastructure in Bryum argenteum Hedw., specimens were cultured for 10 days under four UV‐B treatments (2.75, 3.08, 3.25 and 3.41 W m^–2), simulating depletion of 0% (control), 6%, 9% and 12% of stratospheric ozone at the latitude of Shapotou, a temperate desert area of northwest China. Analyses showed malondialdehyde content significantly increased, whereas chlorophyll (Chl) fluorescence parameters and Chl contents decreased with increased UV‐B intensity. These results corresponded with changes in thylakoid protein complexes and chloroplast ultrastructure. Overall, enhanced UV‐B radiation leads to significant decreases in photosynthetic function and serious destruction of the chloroplast ultrastructure of B. argenteum. The degree of negative influences increased with the intensity of UV‐B radiation. These results may not only provide a potential mechanism for supplemental UV‐B effects on photosynthesis of moss crust, but also establish a theoretical basis for further studies of adaptation and response mechanisms of desert ecosystems under future ozone depletion.     

Exogenous application of IAA alleviates effects of supplemental ultraviolet‐B radiation in the medicinal plant Withania somnifera Dunal

• Supplemental (s)‐UV‐B radiation has adverse effects on the majority of plants. The present study was conducted to evaluate the effects of exogenous application of the growth hormone indole acetic acid (IAA) on various morphological, physiological and biochemical characteristics of Withania somnifera, an indigenous medicinal plant, subjected to s‐UV‐B.
• The s‐UV‐B‐treated plants received ambient + 3.6 kJm^?2·day^?1 biologically effective UV‐B, and IAA was applied at two doses (200 and 400 ppm) to s‐UV‐B‐exposed plants.
• The plant was forced to compromise its growth, development and photosynthetic patterns to survive under s‐UV‐B by increasing concentrations of secondary metabolites and antioxidants (thiol, proline, ascorbic acid, α‐tocopherol, ascorbate peroxidase, catalase, glutathione reductase, peroxidase, polyphenol oxidase, superoxide dismutase) to counteract oxidative stress. Increases in secondary metabolites were evidenced as increased activity of phenylpropanoid pathway enzymes: phenylalanine ammonia lyase, cinnamyl alcohol dehydrogenase, 4‐coumarate CoA ligase, chalcone isomerase and dihydroflavonol reductase. Application of different IAA doses reversed the detrimental effects of s‐UV‐B on W. somnifera by improving growth and photosynthesis and reducing concentrations of secondary metabolites and non‐enzymatic antioxidants. Antioxidant enzymes, however, had a synergistic effect on s‐UV‐B treatment and IAA application.
• The effects of s‐UV‐B on W. somnifera are ameliorated to varying degrees upon exogenous IAA application, and synergistic enhancement of antioxidant enzymes under s‐UV‐B+IAA treatment might be responsible for the partial recuperation of growth and plant protein content, as a UV‐B‐exposed plant is forced to allocate most of its photosynthate towards production of enzymes related to antioxidant defence.

     

Effects of enhanced UV-B radiation on plant physiology and growth on the Tibetan Plateau: a meta-analysis

Uncertainties about the response of plant physiology and growth to enhanced UV-B radiation cause uncertainty to predict how plant production will vary under future radiation change on the Tibetan Plateau. Here, we used a meta-analysis approach to test the influence of UV-B radiation on plant physiology and growth. This hypothesis was tested by investigating the response of plants, which was expressed by some measurable variables. Enhanced UV-B radiation decreased plant biomass, plant height, basal diameter, leaf area index, maximal PSII efficiency, and Chl a+b, but increased intercellular CO₂ concentration, malondialdehyde (MDA), hydrogen peroxide, superoxide anion radical, peroxidase, ascorbate peroxidase, proline and UV-B absorbing compounds. The effect of enhanced UV-B radiation on net photosynthesis rate (P _n ) increased with mean annual precipitation and experimental duration. The effect of enhanced UV-B radiation on MDA decreased with experimental duration. The effect of enhanced UV-B radiation on superoxide dismutase (SOD) increased with the magnitude of enhanced UV-B radiation. Forests rather than grasslands exhibited a positive response of SOD and a negative response of P _n to enhanced UV-B radiation. Therefore, the effect of enhanced UV-B radiation on alpine plants varied with ecosystem types. Local climate conditions may regulate effects of enhanced UV-B radiation on alpine plants.     

紫外线-B辐射与3种植物幼苗的光合作用——光合作用对紫外线-B敏感性的比较

紫外线 - B辐射降低豌豆、大豆和黄瓜 3种植物幼苗的净光合作用速率 ( Pn)和表观量子效率 ( AQE) ,Pn和 AQE的减少幅度随紫外线 - B辐射时间的延长逐渐增大。 3种植物的光合作用对紫外线 - B辐射的敏感性依次为 :豌豆 >黄瓜 >大豆 ,光合作用光抑制的发生程度与光合作用对 UV- B辐射敏感性有内在联系。定量分析显示 ,光合作用光抑制的发生程度随UV- B辐射时间增加 ,是 UV- B辐射剂量的累积效应。     

UV‐B impact on aphid performance mediated by plant quality and plant changes induced by aphids

Plants face various abiotic and biotic environmental factors and therefore need to adjust their phenotypic traits on several levels. UV‐B radiation is believed to impact herbivorous insects via host plant changes. Plant responses to abiotic challenges (UV‐B radiation) and their interaction with two aphid species were explored in a multifactor approach. Broccoli plants [Brassica oleracea L. convar. botrytis (L.), Brassicaceae] were grown in two differently covered greenhouses, transmitting either 80% (high UV‐B) or 4% (low UV‐B) of ambient UV‐B. Three‐week‐old plants were infested with either specialist cabbage aphids [Brevicoryne brassicae (L.), Sternorrhyncha, Aphididae] or generalist green peach aphids [Myzus persicae (Sulzer), Sternorrhyncha, Aphididae]. Plants grown under high‐UV‐B intensities were smaller and had higher flavonoid concentrations. Furthermore, these plants had reduced cuticular wax coverage, whereas amino acid concentrations of the phloem sap were little influenced by different UV‐B intensities. Cabbage aphids reproduced less on plants grown under high UV‐B than on plants grown under low UV‐B, whereas reproduction of green peach aphids in both plant light sources was equally poor. These results are likely related to the different specialisation‐dependent sensitivities of the two species. The aphids also affected plant chemistry. High numbers of cabbage aphid progeny on low‐UV‐B plants led to decreased indolyl glucosinolate concentrations. The induced change in these glucosinolates may depend on an infestation threshold. UV‐B radiation considerably impacts plant traits and subsequently affects specialist phloem‐feeding aphids, whereas aphid growth forces broccoli to generate specific defence responses.     

Variation of the enhanced biologically damaging solar UV due to clouds.

The variation of the biologically damaging solar UV (UVBE) enhanced by clouds above that of clear sky UVBE has been investigated. This was undertaken for summer through to winter for SZA of 5 to 60 degrees employing an integrated automatic cloud and spectral UV measurement system that recorded the solar UV spectra and the sky images at five minute intervals. The UVBE calculated with action spectra with higher relative effectiveness in the UVA produced the lower percentage of cloud enhanced cases. The DNA UVBE provided the highest percentage of cloud enhanced cases compared to the total number of UV scans with 2.2% cloud enhanced cases. As a comparison, the plant and fish melanoma UVBE provided the lowest percentage of cloud enhanced cases with 0.6 to 0.8% cloud enhanced cases. For the cases of cloud enhanced UVBE, the average ratio of the measured UVBE to calculated cloud free UVBE for the photokeratitis, cataracts, plant, generalized plant damage and fish melanoma action spectra was 1.21 to 1.25. In comparison, the highest value of 1.4 was for the DNA action spectrum.     

UV-B辐射增强对陆地植物次生代谢的影响

平流层臭氧的减薄已导致地表中波紫外辐射(UV-B,280～320nm)增强,由于UV-B能被许多生物大分子如蛋白质和核酸吸收并引起分子构象的变化,因此可对植物的各方面产生影响。本文将近年来特别是近5年的UV-B辐射增强对植物次生代谢物影响的研究工作进行了综述。主要包括：UV-B辐射增强对植物紫外吸收物的影响和可能的机制;环境因子的复合作用对植物紫外吸收物的影响和可能的机制;UV-B辐射增强对次生代谢物影响的生态学意义。并对该领域未来的研究作了展望。